The genetic revolution[edit]

The genetic revolution in studies of human evolution started when Vincent Sarich and Allan
Wilson measured the strength of immunological cross-reactions of blood serum albumin between
pairs of creatures, including humans and African apes (chimpanzees and gorillas).
[20]
The strength of
the reaction could be expressed numerically as an immunological distance, which was in turn
proportional to the number of amino acid differences between homologous proteins in different
species. By constructing a calibration curve of the ID of species' pairs with known divergence times
in the fossil record, the data could be used as a molecular clock to estimate the times of divergence
of pairs with poorer or unknown fossil records.
In their seminal 1967 paper in Science, Sarich and Wilson estimated the divergence time of humans
and apes as four to five million years ago,
[20]
at a time when standard interpretations of the fossil
record gave this divergence as at least 10 to as much as 30 million years. Subsequent fossil
discoveries, notably Lucy, and reinterpretation of older fossil materials, notably Ramapithecus,
showed the younger estimates to be correct and validated the albumin method. Application of
the molecular clock principle revolutionized the study of molecular evolution.
The quest for the earliest hominin[edit]
In the 1990s, several teams of paleoanthropologists were working throughout Africa looking for
evidence of the earliest divergence of the Hominin lineage from the great apes. In 1994, Meave
Leakey discovered Australopithecus anamensis. The find was overshadowed by Tim White's 1995
discovery of Ardipithecus ramidus, which pushed back the fossil record to 4.2 million years ago.
In 2000, Martin Pickford and Brigitte Senut discovered in the Tugen Hills of Kenya a 6-million-year-
old bipedal hominin which they named Orrorin tugenensis. And in 2001, a team led by Michel
Brunet discovered the skull of Sahelanthropus tchadensis which was dated as 7.2 million years ago,
and which Brunet argued was a bipedal, and therefore a hominid.
Human dispersal[edit]

A model of human migration, based from divergence of themitochondrial DNA (which indicates
the matrilineage).
[21]
Timescale (kya) indicated by colours.

A "trellis" (as Wolpoff called it) that emphasizes back-and-forth gene flow among geographic regions.
[22]

Different models for the beginning of the present human species.
See also: Early human migrations, Recent African origin of modern humans and Multiregional
hypothesis
Anthropologists in the 1980s were divided regarding some details of reproductive barriers and
migratory dispersals of the Homo genus. Subsequently, genetics has been used to investigate and
resolve these issues. According to the Sahara pump theory evidence suggests that
genus Homo have migrated out of Africa at least three times (e.g. Homo erectus, Homo
heidelbergensis and Homo sapiens).
The Out-of-Africa model proposed that modern H. sapiens speciated in Africa recently (approx.
200,000 years ago) and the subsequent migration through Eurasia resulted in nearly complete
replacement of otherHomo species. This model has been developed by Chris Stringer and Peter
Andrews.
[23][24]
In contrast, themultiregional hypothesis proposed that Homo genus contained only a
single interconnected population as it does today (not separate species), and that its evolution took
place worldwide continuously over the last couple million years. This model was proposed in 1988
by Milford H. Wolpoff.
[25][26]

Progress in DNA sequencing, specifically mitochondrial DNA (mtDNA) and then Y-chromosome
DNAadvanced the understanding of human origins.
[27][28][29]
Sequencing mtDNA and Y-DNA sampled
from a wide range of indigenous populations revealed ancestral information relating to both male
and female genetic heritage.
[30]
Aligned in genetic tree differences were interpreted as supportive of
a recent single origin.
[31]
Analyses have shown a greater diversity of DNA patterns throughout Africa,
consistent with the idea that Africa is the ancestral home of mitochondrial Eve and Y-chromosomal
Adam.
[32]

Out of Africa has gained support from research using female mitochondrial DNA (mtDNA) and the
male Y chromosome. After analysing genealogy trees constructed using 133 types of mtDNA,
researchers concluded that all were descended from a female African progenitor,
dubbed Mitochondrial Eve. Out of Africa is also supported by the fact that mitochondrial genetic
diversity is highest among African populations.
[33]

A broad study of African genetic diversity, headed by Sarah Tishkoff, found the San people had the
greatest genetic diversity among the 113 distinct populations sampled, making them one of 14
"ancestral population clusters". The research also located the origin of modern human migration in
south-western Africa, near the coastal border of Namibia andAngola.
[34]
The fossil evidence was
insufficient for Richard Leakey to resolve this debate.
[35]
Studies of haplogroups in Y-chromosomal
DNA and mitochondrial DNA have largely supported a recent African origin.
[36]
Evidence from
autosomal DNA also predominantly supports a Recent African origin. However evidence
for archaic admixture in modern humans had been suggested by some studies.
[37]

Recent sequencing of Neanderthal
[38]
and Denisovan
[39]
genomes shows that some admixture
occurred. Modern humans outside Africa have 2-4% Neanderthal alleles in their genome, and
some Melanesians have an additional 4-6% of Denisovan alleles. These new results do not
contradict the Out of Africa model, except in its strictest interpretation. After recovery from a genetic
bottleneck that might be due to the Toba supervolcano catastrophe, a fairly small group left Africa
and briefly interbred with Neanderthals, probably in the middle-east or even North Africa before their
departure. Their still predominantly African descendants spread to populate the world. A fraction in
turn interbred with Denisovans, probably in south-east Asia, before populating
Melanesia.
[40]
HLA haplotypes of Neanderthal and Denisova origin have been identified in modern
Eurasian and Oceanian populations.
[14]

There are still differing theories on whether there was a single exodus or several. A multiple
dispersal model involves the Southern Dispersal theory,
[41]
which has gained support in recent years
from genetic, linguistic and archaeological evidence. In this theory, there was a coastal dispersal of
modern humans from the Horn of Africa around 70,000 years ago. This group helped to populate
Southeast Asia and Oceania, explaining the discovery of early human sites in these areas much
earlier than those in the Levant.
[41]

A second wave of humans may have dispersed across the Sinai peninsula into Asia, resulting in the
bulk of human population for Eurasia. This second group possibly possessed a more sophisticated
tool technology and was less dependent on coastal food sources than the original group. Much of
the evidence for the first group's expansion would have been destroyed by the rising sea levels at
the end of each glacial maximum.
[41]
The multiple dispersal model is contradicted by studies
indicating that the populations of Eurasia and the populations of Southeast Asia and Oceania are all
descended from the same mitochondrial DNA lineages, which support a single migration out of
Africa that gave rise to all non-African populations.
[42]